KARYOLOGY OF SPECIES OF POLYOMMATUS (AGRODIAETUS) FROM TURKEY: NEW DATA AND THEIR TAXONOMIC CONSEQUENCES (LEPIDOPTERA: LYCAENIDAE)

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1 Vol.5 No LUKHTANOV et al.: Polyommatus Karyology in Turkey 1 HOLARCTIC LEPIDOPTERA, 5(1): 1-8 KARYOLOGY OF SPECIES OF POLYOMMATUS (AGRODIAETUS) FROM TURKEY: NEW DATA AND THEIR TAXONOMIC CONSEQUENCES (LEPIDOPTERA: LYCAENIDAE) VLADIMIR A. LUKHTANOV1, NIKOLAI P. KANDUL1, WILLY O. DE PRINS2 AND DIRK VAN DER POORTEN3 1 Dept. of Entomology, Faculty of Biology, University of St. Petersburg, Universitet. nab. 7/9, RU St. Petersburg, Russia 2 Diksmuidelaan 176, B-2600 Antwerp, Belgium 3 Lanteernhofstraat 26, B-2140 Antwerp, Belgium ABSTRACT.- The karyotypes of the following taxa of the subgenus Polyommatus (Agrodiaetus) from Turkey were investigated: P. (A.) alcestis (Zerny, 1932) (n=19), P. (A.) demavendi (Pfeiffer, 1938) (n=ca ), P. (A.) antidolus (Rebel, 1901) (n=39-41), P. (A.) hopfferi (Herrich-Schaffer, [1851]) (n=15), P. (A.) kurdistanicus (Forster, 1961) (n=ca ), P. (A.) sp. (? huberti Carbonell, 1993) (n=22), P. (A.) merhaba De Prins et al., 1991 (n=16-17), P. (A.) aserbeidschanus turcicola (Kocak, 1977) (n=19-20), P. (A.) elbursicus zapvadi (Carbonell, 1993) (n=17-18), P. (A.) firdussii pseudactis (Forster, 1960) (n=22-25), P. (A.) cyaneus (Staudinger, 1899) (), P. (A.) mrcicus (Kocak, 1977) () and P. (A.) iphigenia (Herrich-Schaffer, [1851]) (n=13). The taxonomic position of the investigated taxa is discussed according to their karyotypes. ZUSAMMENFASSUNG.- Die Karyotypen der folgenden Taxa der Untergattung Polyommatus (Agrodiaetus) aus der Tiirkei wurden untersucht: P. (A.) alcestis (Zerny, 1932) (n=19), P. (A.) demavendi (Pfeiffer, 1938) (n=ca ), P. (A.) antidolus (Rebel, 1901) (n=39-41), P. {A.) hopfferi (Herrich-Schaffer, [1851]) (n=15), P. (A.) kurdistanicus (Forster, 1961) (n=ca ), P. (A.) sp. (? huberti Carbonell, 1993) (n=22), P. (A.) merhaba De Prins et al., 1991 (n=16-17), P. (A.) aserbeidschanus turcicola (Kocak, 1977) (n=19-20), P. (A.) elbursicus zapvadi (Carbonell, 1993) (n=17-18), P. (A.) firdussii pseudactis (Forster, 1960) (n=22-25), P. (A.) cyaneus (Staudinger, 1899) (), P. (A.) turcicus (Ko9ak, 1977) () und P. (A.) iphigenia (Herrich-Schaffer, [1851]) (n=13). Die taxonomische Stellung der untersuchten Taxa wird Ihren Karyotypen entsprechend diskutiert. KEY WORDS: Armenia, Asia Minor, chromosomes, distribution, genetics, Iran, Lebanon, Middle East, Near East, Palearctic, Rhopalocera, spermatogenesis, systematics, variation, taxonomy. The diverse Polyommatus subgenus Agrodiaetus Hiibner, [1822], Many species of the subgenus Agrodiaetus occur in Turkey: is composed of several species, subspecies and forms, of which the Hesselbarth, van Oorschot and Wagener (1995) list 35 species, exact taxonomic position is not always clear. Consequently, Agrodi- Fortunately, the karyotypes of many taxa in this region have been aetus is one of the most difficult groups of Polyommatus among studied by de Lesse (1959a,b,c; 1960a,b,c; 1961; 1962; 1963b,c). Palaearctic Rhopalocera. In some cases it is even impossible to Two species (P. (A.) poseidon and P. (A.) theresiae) have been separate sympatric species, because morphological differences studied by Kandul and Lukhtanov (1997). amongst them are difficult to recognise. It is also difficult to Our own karyological studies have two aims. The first one is to understand whether allopatric populations belong to separate species establish the chromosome number of recently described taxa from or whether they constitute local forms of a species living in a wide Turkey; the second one is to study the taxonomic relationship area. In many cases, it is easy enough to describe such populations through karyological analysis in some difficult species complexes: as subspecies, since most of them show constant differences and are demavendi - alcestis, antidolus - kurdistanicus and ninae - elbursicus geographically isolated, but at the same time it is virtually impossible - turcicola. In the light of the new publication of Eckweiler and to unite them into polytypic species using only morphological Ha'user (1997), we would like to discuss the taxonomic position of characters. P. (A.) theresiae, the karyotype of which was studied earlier (Kandul While external morphological characters in this subgenus are and Lukhtanov, 1997). homogeneous, the karyotypes show many dissimilarities. The number of chromosomes can vary from n=8 in P. (A.) nephohiptamenos MATERIAL AND METHODS (Brown and Coutsis, 1978) to n= in P. (A.) dolus (Hlibner,,:,,, 18231) (de Lesse. 1961) in^ix and j this,unumber u is specific -c in most cases. O^y males were used to establish the chromosome numbers..,,.... because the karyological study of females is a more difficult A complete list of known chromosome numbers in the subgenus,j,...,,,tt.,,, problem. The abdomens, or only the testes, of living, fresh (not Aerodtaetus was recently published (Hesselbarth, van Oorschot and., -, -, -, Wagener, irir.c, 1995). Although..,, some species have, identical.,,., or only, worn) adult.. specimens, were placed in small vials in which a freshly K, _... n /, j / mixed solution of 3 parts 96% ethanol and 1 part 100% acetic acid slightly different chromosome numbers (e.g., P. A.) damone simrica....«,... 1onn- n /. _c innnx\u was kept. The individual fixations were given a code number which (Staudinger, 1899), and P. (A.) morgam (Le Cerf, 1909)), the 6 ', was also noted on the paper in which the donor butterfly was kept, structure of their karyotypes vanes, i.e., in the number of macrochro- _,....J..,r>orv»>r ; inr,cn A The vials were immediately put into a thermos filled with icy water mosomes (Lukhtanov, 1989; Munguira et al., 1995). As can be. / b... /. r ~. in order to keep the fixations at a low temperature of 0-4 C. The concluded from the cited work of de Lesse (1961), it is absolutely.. «-,,»,,,... : whole operation was done in the field. After the expedition, the vials necessarv (but not always enough) to examine the karyotypes or all,,......,.,,,. were put into a regular refngerator at 4 C, where they could be kept the seographic forms in this subgenus in order to understand their. e. B. J for a long period. During the subsequent cytological studies, squash taxonomic position.

2 r 2 LUKHTANOV el al.: Polyommatus Karyology in Turkey HOLARCTIC LEPIDOPTERA TABLE 1. Chromosome numbers of the taxa of the subgenus Polyommatus (Agrodiaetus Hiibner) examined from Turkey Taxon, locality, collecting date Specimen Codes Chromosome Number Number and stage of examined cells P. (A.) alcestis (Zerny, 1932) Hakkari: Dez Valley, 1500m 19 July P. (A), demavendi (Pfeiffer, 1938) Hakkari: Dez Valley, 1500 m 19 July ib ibid ib Hakkari: 16 km N Bagishli, 2000m 26 July Bitlis: Kuzgunkiran Gecidi, m 26 July ib ib ib P. (A.) antidolus (Rebel, 1901) Hakkari: 16 km N Bagishli, 2000m 6Augl ib ib ib ib ib ib ib ib ib Van: 32 km N Catak, m 10 Aug P. (A.) hopfferi (Herrich-Schaffer, [1851]) Hakkari: Dez Valley, m 6 Aug ib P. (A.) kurdistanicus (Forster, 1961) Van: 32 km N Catak, m 10 Aug P. (A.) sp. (? huberti Carbonell, 1993) Van: Guzeldere Gecidi, m 4-5 Aug ib ib P. (A.) merhaba De Prins et al Artvin: 10 km SW Yusufeli, 900m 15 July ib P. (A.) aserbeidschanus turcicola (Koc.ak, 1977) Van: 32 km N Catak, m July ib ib Van: Zernek Baraji, m 29 July-2 Aug ib Van: Guzeldere Gecidi, m 4-5 Aug P. (A.) elbursicus zapvadi (Carbonell, 1993) Van: Zernek Baraji, m 29 July-2 Aug ib ib ib n=19 n=ca. 70 n=39 n=ca n=41 n=40 n=ca n=41 n=40+m n=40 n=ca n=41 n=ca n=15 n=15 n=ca n=ca n=22 n=22 n=16 n=17 n=19 n=17 n= M-I 5 M-I 7 M-l 5 M-I 4 M-I 8 M-I 9 M-I, Pr., 3 Pr 1 17 M-I 1, 2 Pr. 24 M-I, 1 Pr 7 M-I M-I 9 M-I, 9 M-I Pr.

3 Vol. 5 No LUKHTANOV et al.: Polyommatus Karyology in Turkey 3 P. (A.) firdussii pseudactis (Forster, 1960) Bitlis: Kuzgunkiran Gecidi, m 26 July ' ib Van: 32 km NE Catak, m 28 July Van: Guzeldere Gecidi, m 4-5 Aug ib ib P. (A.) cyaneus (Staudinger, 1899) Van: Zernek Baraji, m 29 July-2 Aug ib ib ib ib P. (A.) turcicus (Ko^ak, 1977) Erzurum: 30 km NNE Erzurum, m July Van: 32 km NNE Catak, m July ib ib ib ib ib P. (A.) iphigenia (Herrich-Schaffer, [1851]) Bitlis: Kuzgunkiran Ge?idi, m 26 July Van: 32 km NNE Catak, m July n=25 n=25+m n=25 n=22 n=23 n=ca n=17+m n=16+2m n=ca. 24 2n=ca n=13 n=13 8 M-I 8 M-I 15 M-I 4 M-I 7 M-I, 5 M-I 3 Mit. 4Pr. 2Pr. 2 and M-II 1 preparations of the testes were made, after being stained with aceticorcein. Haploid chromosome numbers (n) were determined in metaphase-i (M-I) of spermatogenesis, more rarely also in metaphase-ii (M-II), the prophase of the first meiotic division (Pr.), or the metaphase of gonadial mitosis (Mit.) Negatives and photographs of the studied metaphase preparations are kept in the Department of Entomology, University of St. Petersburg, Russia. The set specimens of the donor butterflies are kept in the collections of the Institute of Systematics and Population Biology (Zoological Museum), Amsterdam, the Netherlands, and the Flemish Lepidoptera Collection, Antwerp, Belgium. RESULTS The results of the karyological studies are summarized in Table 1 and in Fig. 1-15, with comments below. The taxa are listed (slightly modified) according to Eckweiler and Hauser (1997). P. (A.) alcestis (Zerny, 1932) (Fig. 1). In metaphase-i, all bivalent chromosomes form a gradient series. The karyotype contains no exceptionally big or small bivalents. P. (A.) demavendi (Pfeiffer, 1938) (Fig. 2). The chromosome numbers were only approximately established. They are similar in several examined populations (Table 1). The karyotype contains 2 very large and 2 median bivalents. All other bivalents are relatively small and form a gradient series in metaphase-i. P. (A.) antidolus (Rebel, 1901) (Fig. 3). The number of chromosomes varies between individuals from n=39 to n=41. The bivalents are differently shaped, ranging from very small to large. In metaphase-i, 5 to 6 large bivalents can be recognized, from which the first is somewhat bigger than the other. P. (A.) hopfferi (Herrich-Schaffer, [1851]) (Fig. 4). The variability in the chromosome numbers was not found in two examined specimens. The bivalents in metaphase-i are large and approximately of the same size. P. (A.) kurdistanicus (Forster, 1961) (Fig. 5). The chromosome number was only approximately determined. The karyotype contains 3 very large bivalents. P. (A.) sp. (? huberti Carbonell, 1993) (Fig. 8). In two examined specimens, no variability of the chromosome number was found. In metaphase-i, all bivalents form a gradient series. P. (A.) merhaba de Prins, van der Poorten, Bone, van Oorschot, Riemis & Coenen, 1991 (Fig. 9-10). One specimen showed an intraindividual variability of n=16 to n=17. In metaphase-i, all bivalents in both karyotype variants (n=16 Fig. 10, n=17 Fig 9) are of the same size. P. (A.) aserbeidschanus turcicola (Ko9ak, 1977) (Fig. 11). The most common chromosome number in this species is. One specimen showed an intraindividual variability of n=19 to. In metaphase-i, all bivalents form a gradient series. P. (A.) elbursicus zapvadi (Carbonell, 1993) An intraindividual variability in the number of chromosomes was found, ranging from to n=19. In metaphase-i, all bivalents form a gradient series. The karyotype shows no extraordinary large or small bivalents. P. (A.) firdussii pseudactis (Forster, 1960) (Fig. 12). In this species, not only variability in the number of bivalents was encountered, but also the absence or presence of a small extra element (m) was not

4 li *..,» ;.*» %* / ' **! ' Jill V-' 5Mf %* \ m 8 Fig Karyotypes of certain Polyommatus (Agrodiaetus) taxa: 1. P. alcestis (Zerny, 1932), specimen Nr 92024, M-I, n=19, Hakkari, Dez valley, 1500m, 19 July 1992; 2. P. demavendi (Pfeiffer, 1938), specimen Nr 92048, M-I,, Bitlis, Kuzgunkiran Gefidi, m, 26 July 1992; 3. P. antidolus (Rebel, 1901), specimen Nr 92124, M-I, n=40, Hakkari, 16 km N Bagishli, 2000m, 6 Aug 1992; 4. P. hopfferi (Herrich-Schaffer, [1851]), specimen Nr 92135, M-I, n=15, Hakkari, Dez valley, m, 6 Aug 1992; 5. P. kurdistanicus (Forster, 1961), specimen Nr 92148, M-I, n=ca , Van, 32 km N Catak, m, 10 Aug 1992; 6. P. poseidon (Herrich-Schaffer, [1851]), specimen Nr 92002, prophase of the first division, n=19, Artvin, 10 km SW Yusufeli, 900m, 15 July 1992 (according to Kandul and Lukhtanov, 1997: fig. 1,1); 7. P. theresiae Schurian, van Oorschot & van den Brink, 1992, specimen Nr 94033, M-I, n=ca , Konya, Taskent, July 1994 (according to Kandul and Lukhtanov, 1997: fig. 1,6); 8. P. sp. (Ihuberti Carbonell, 1993), specimen Nr 92114, M-I, n=22, Van, Guzeldere Gecidi, m, 4-5 Aug 1992.

5 * J 10 f- * :-'. * «* ** 12! * Fig Karyotypes: 9. P. merhaba De Prins el al. 1991, specimen Nr 92001, M-I, n=17, Artvin, 10 km SW Yusufeli, 900m, 15 July 1992; 10. P. merhaba De Prins ei al. 1991, specimen Nr 92001, M-I, n=16, Artvin, 10 km SW Yusufeli, 900m, 15 July 1992; 11. P. aserbeidschanus turcicola (Kocak, 1977), specimen Nr 92081, M-I,, Van, Zernek Baraji, m, 29 July-2 Aug 1992; 12. P. firdussii pseudactis (Forster, 1960), specimen Nr 92043, M-I, n=25, Bitlis, Kuzgunkiran Gecidi, m, 26 July 1992; 13. P. cyaneus (Staudinger, 1899), specimen Nr 92082, M-I,, Van, Zernek Baraji, m, 29 July-2 Aug 1992; 14. P. turcicus (Kocak, 1977), specimen Nr 92019, M-I,, 30 km NNE Erzurum, m, July 1992; 15. P. iphigenia (Herrich-Schaffer, [1851]), specimen Nr 92057, M-I, n=13, Van, 32 km NNE Catak, m, July 1992.

6 6 LUKHTANOV et al.: Polyommatus Karyology in Turkey HOLARCTIC LEPIDOPTERA constant. Furthermore, variability in specimens from different localities is significant. Because from every locality studied, only one butterfly produced suitable preparations in metaphase, we can not conclude at this moment whether this phenomenon is due to geographical or to indiviual variability. The bivalents in metaphase-i are all of approximatively the same medium size. P. (A.) cyaneus (Staudinger, 1899) (Fig. 13). In one specimen the chromosome number was stable, whereas in another one, a variability in the number of bivalents and the presence of a small, extra, probably univalent element (m), was found. In the variant of the karyotype with, all bivalents show a gradient series in metaphase-i. P. (A), turcicus (Ko9ak, 1977) (Fig. 14). The chromosome number of was identical in all examined specimens from two populations. In metaphase-i, all bivalents form a gradient series. P. (A.) iphigenia (Herrich-Schaffer, [1851]) (Fig. 15). The chromosome number of n=13 was identical in both specimens from two populations. All bivalents in metaphase-i and chromosomes (in phase M-II) form a gradient series. DISCUSSION P. (A.) alcestis (Zerny, 1932) Our statements about the number of chromosomes and the structure of the karyotypes completely conform to those of de Lesse (1960a, b). From the studies of de Lesse (1960a,b), Larsen (1975) and our data, it is clear that the populations of P. (A.) alcestis can be divided into two groups with different chromosome numbers. The western group has a stable chromosome number of and contains the populations from Turkey (except southeastern Turkey) and Lebanon. Only in very few specimens does the number n=21 occur (de Lesse, 1960b). As can be seen from the figures given by de Lesse (1960b: Fig. 5c,d), this fact can be ascribed to a very small extra element, which occurs at the edge of the metaphase plate and can best be regarded as a B-chromosome. The oriental group also has a stable chromosome number of n=19 and includes the populations of Iran. The only specimen we examined from southeastern Turkey (Hakkari) belongs to the oriental subspecies, according to its chromosome number. This is not surprising, since in southeastern Turkey more Iranian taxa are found which do not occur further to the west or the north in the other Turkish provinces (cf. Hesselbarth, van Oorschot and Wagener, 1995). P. (A.) demavendi (Pfeiffer, 1938) Our statements about the number of chromosomes and the structure of the karyotypes completely agree with those of de Lesse (1960a,b; 1963a), who studied butterflies from Iran and Turkey. We found the same karyotype in several areas of the Transcaucasus: in Azerbaijan (Nakhichevan) and Armenia (Lukhtanov, unpubl.). This demonstrates the wide distribution of the species in the Near East. P. (A.) antidolus (Rebel, 1901) P. (A.) kurdistanicus (Forster, 1961) These species belong, according to Eckweiler and Hauser (1997), to a group of closely related taxa: P. (A.) antidolus (Rebel, 1901), P. (A.) kurdistanicus (Forster, 1960), P. (A.) morgani (Le Cerf, 1909) and P. (A.) femininoides (Eckweiler, 1987). According to de Lesse (1960a, 1961), this group contains at least three different species: sp. 1 with n=60-62 (Turkey: Van), sp. 2 with n=42 (Turkey: Pertek), and sp. 3 with n=25-26 (Iran: Sanandaj). During our studies of this group, we found two variants of the karyological types: n=ca and n=39-41, both encountered in the region of Lake Van. These karyotypes are very similar, and probably identical to those of "sp. 1" and "sp. 2" of de Lesse in regard to the number of chromosomes and the number of large bivalents. Wagener and his colleagues (Hesselbarth, van Oorschot & Wagener, 1995) ascribed these three karyotypes to the following taxa: kurdistanicus (n=60-62), antidolus (n=42) and morgani (n=25-26). This looks logical but needs confirmation, in the first place by examining the karyotypes of kurdistanicus and morgani from the type-localities. The first author examined the specimens in the Museum National d'histoire Naturelle, Paris, for which de Lesse established n=25-26, and found out that they differed considerably in characters of external morphology (more blue and darker scaling) from the type-specimens of morgani. Furthermore, they were collected far to the north of the type-locality of morgani. P. (A.) hopfferi (Herrich-Schaffer, [1851] P. (A.) iphigenia (Herrich-Schaffer, [1851]). Our statements about the karyotypes of these two species conform with the studies of de Lesse (1959a,b,c; 1960a,c). P. (A.) theresiae Schurian, van Oorschot and van den Brink, 1992 This species was recently separated from the similar, sympatric P. (A.) poseidon (Schurian et al., 1992). Our statements about the number of chromosomes confirm that P. (A.) theresiae (Fig. 7) and P. (A.) poseidon (Fig. 6) are not conspecific (Kandul and Lukhtanov, 1997). In Hesselbarth, van Oorschot and Wagener (1995). this species was placed in the group of transcaspicus and elbursicus. However, according to the recent study of Eckweiler and Hauser (1997), this taxon belongs to the group of P. (A.) dama (Staudinger, 1892). P. (A.) theresiae can be separated from P. (A.) dama by the presence of a white stripe on the underside of the hindwings, by its brighter blue color of the upperside, and by the presence of a concentrated androconial patch on the upperside of the forewing, which is absent in P. (A.) dama dama. P. (A.) theresiae shows no significant karyological differences to P. (A.) dama, of which the chromosome number was established by de Lesse (1959c) as n=41-42 (Malatya). Both taxa are probably allopatric. According to Hesselbarth, van Oorschot and Wagener, 1995), both occur in the "Umgebung Kahramanmaras, m: M.-E.VII.1929". But this information is based upon old material. Only further studies can establish whether P. (A.) theresiae and P. (A.) dama should be considered as two subspecies of the same species or as two separate species. Our statements about the chromosome number of P. (A.) theresiae furthermore prove that the recently described taxon P. (A.) larseni (Carbonell, 1994), with n=25-26 (see Larsen, 1975) cannot be considered as a subspecies of P. (A.) theresiae. According to the chromosome number and the color and shape of the wings, it belongs to the group of P. damocles (Herrich-Schaffer, [1844]). The members of the latter group have a similar chromosome number (-26) and a similar karyotype structure (Lukhtanov et al., 1997). P. (A.) sp. (Ihuberti Carbonell, 1993) Carbonell (1993) designated a specimen as the holotype of this taxon from a series from Agri, which was determined by de Lesse (1960a, 1963c) as "'transcaspica ninae" and for which chromosome numbers n=33 and n=34 were established. Two.specimens from the Giizeldere Ge9idi (Van) that were studied, which were mentioned in Hesselbarth, van Oorschot and Wagener (1995) as P (A.) huberti, have n=22. Probably, we can conclude from this result that two different taxa are involved under the name P. (A.) huberti.

7 Vol. 5 No LUKHTANOV et al.: Polyommatus Karyology in Turkey 7 P. (A.) merhaba De Prins et al., 1991 P. (A.) cyaneus (Staudinger, 1899) Our statements on the chromosome number of P. (A.) cyaneus completely agree with those of de Lesse (1963b), taken from the populations of Lake Van. Typical for this species is a significant geographical variability in the chromosome numbers ranging from n=16-17 to n=22 (Lukhtanov, 1989). P. (A.) merhaba, which is taxonomically very close to and as far as known allopatric with P. (A.) cyaneus, has n= Because the chromosome number of P. (A.) merhaba falls within the known range of P. (A.) cyaneus, we cannot propose any definitive statements about the taxonomic position of P. (A.) merhaba (species or subspecies) from the karyological point of view. The morphological differences, however, are clear: P. (A.) merhaba is of a brighter blue color and it lacks the black venation in the marginal area of the upperside. P. (A.) aserbeidschanus turcicola (Ko ak, 1977) P. (A.) elbursicus lapvadi (Carbonell, 1993). Both of these taxa belong to the P. (A.) transcaspicus (Heyne, [1895]) group. In earlier studies, this group was divided into three subgroups (Lukhtanov, 1989): an oriental subgroup (P. transcaspicus sensu Lukhtanov, 1989) with n=52-53, a central subgroup. P. aserbeidschanus sensu Lukhtanov, 1989) with chromosome numbers ranging from n=16 to n=22 in different populations, and an occidental subgroup (P. ninae sensu Lukhtanov, 1989) with n= Each of these subgroups was given specific status, though it was supposed that the central subgroup was heterogeneous and contained several separate species. Field studies during the last few years showed that P. (A.) aserbeidschanus (Forster, 1956) and P. (A.) ninae (Forster, 1956) (Dantchenko, pers. comm.; see also Hesselbarth, van Oorschot and Wagener, 1995: 737) therefore should be considered specifically distinct. Subsequent studies also confirm that the central subgroup consists of several species. So, Carbonell (1993) showed that the taxon from Van with, and determined by de Lesse (1963c) as P. (A.) poseidon, morphologically and karyologically belongs to P. (A.) elbursicus (Forster, 1956) and occurs sympatrically with P. (A.) aserbeidschanus turcicola. This proves that these taxa of the central subgroup are not conspecific. Our current karyological studies confirm and verify the conclusion that in the region of Lake Van, two karyological and morphological distinguishable taxa of the P. aserbeidschanus complex occur: P. (A.) turcicola (Kogak, 1977) with n=19-20 and P. (A.) elbursicus zapvadi (Carbonell, 1993) with n= P. (A), firdussii pseudactis (Forster, 1960) Our karyological statements principally conform to those of de Lesse (1960a, 1962), who found chromosome numbers for this subspecies in different localities ranging from to n=30. Only one male in our material showed n= P. (A.) turcicus (Kocak, 1977) The karyotype of this species was previously not known, but we were able to establish it. It is interesting from two aspects. First, the karyotype is very stable and shows no variation at all in the number of chromosomes, a condition which is known only from a few species with a wide distribution range, e.g. P. (A.) ripartii (Freyer, 1830), P. (A.) menalcas (Freyer, [1839]) and P. (A.) daman ([Denis & Schiffermiiller], 1775) (see Hesselbarth, van P. Oorschot and Wagener, 1995). Furthermore, P. (A.) turcicus is the only species in the subgenus Agrodiaetus with the chromosome number of, which is the mean and probably ancestral number for all non-tropical groups of Lycaenidae (Lukhtanov, 1993). If this number is a plesiomorphic character state and not evolved secondarily, P. (A.) turcicus can be considered as a primitive species in Agrodiaetus. ACKNOWLEDGEMENTS The authors would like to thank the following colleagues: Dr. J.-P. Bone, F. Carbonell, A. Dantchenko, Dr. W. Eckweiler, H. van Oorschot and Dr. P. S. Wagener, for the fruitful discussions about taxonomical and karyological problems within the subgenus Agrodiaetus; Dr. W. Dierl (Zoologische Staatssammlung Munchen), Prof. Dr. G. Bernard! and Dr. M. J. Pierre (Museum National d'histoire Naturelle, Paris), for the opportunity to study the types.of the taxa described by Dr. W. Forster and H. de Lesse, and the specimens of which the chromosomes were studied by H. de Lesse. The first two authors furthermore would like to thank the Russian Fund for Pure Research (Grant Nr ), for financial support. REFERENCES Brown, J., and J. G. Coutsis Two newly discovered lycaenid butterflies (Lepidoptera, Lycaenidae) from Greece, with notes on allied species. Em. Gaz. (Wallingford), 29: Carbonell, F Contribution a la connaissance du genre Agrodiaetus Hiibner (1822): le complexe ultraspecifique d'a. transcaspica Staudinger, (1899) (Lepidoptera: Lycaenidae). Linn. Belg. 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